Examination of the type materials ofNavicula subtilissima Cleve (Bacillariophyceae)

Navicula subtilissima was described by Cleve as a new species from ‘a dry rivulet at the shore of Imandra, at the foot of Chibinä, Lapland’. However, the examination of the type slides and the unmounted original material using transmission and scanning electron microscopy revealed that there are at least three finely striated similar taxa distinguishable at the specific level. From these taxa, application of the nameN. subtilissima Cleve is established by selecting a lectotype; a second one is described as a new species,N. parasubtilissima, and the third is identified asN. subtilissima var.micropunctata Germain. The last one warrants a specific status by its fine structural differences in having continuous striae consisting of a single elongate areola occluded by a hymen with perforations arranging hexagonally. The occurrence of these three species is shown in Japanese collections.     

Climatic response of budburst in the mountain birch at two areas in northern Fennoscandia and possible responses to global change

The relationship between the climate and budburst of the mountain birch was evaluated for two areas in subarctic (ca 69°N) Fennoscandia; at Abisko, Swedish Lapland, and at Kevo, Finnish Lapland. Thermal time (TT, degree‐day sums) to budburst was calculated for experimental conditions in the laboratory and for in situ observations of budburst. Two types of models predicting leaf emergence in situ were used: 1) TT to budburst for different threshold temperatures based on daily mean (TT_Mean) or daily maximum (TT_Max) temperatures and 2) ecophysiological budburst models. The obtained models were used to estimate effects of a changed climate.
Laboratory experiments of TT to budburst indicated no differences in the thermal requirements at the two areas. Temperature requirements of budburst declined successively during the progression of spring, from ca 250 degree‐days (>+2°C) in January to ca 100 in May. No significant trend in the date of budburst was found over the last 70 (Abisko) or 20 (Kevo) yr. There were some differences in the type of model that best explained the date of budburst in situ at the two areas. For Kevo the best prediction (minimum root mean square error, RMSE) of budburst was obtained by a simple thermal time model (TT_Mean>5.5°C) from 1 January (RMSE=2.1). For budburst at Abisko, models based on daily maximum temperature fitted better than those based on daily means. For Abisko, models based on thermal time accumulation only showed systematic errors in the predicted budburst that were correlated with budburst previous year (BB_PY). Including this apparent memory effect in the model decreased the error by 2.4 d. The best prediction for Abisko was thus obtained using TT_MAX>6.5 (RMSE=3.1) from 1 January.
Using these models to predict the effect of a changed air temperature climate indicate 3–8 d earlier budburst for a one‐degree increase in temperature, the effect being smaller for Kevo than for Abisko. For both areas a change in May temperature has a larger effect on the date of budburst than changes in any other month.     

Ecology of a steppe-tundra gradient in interior Alaska

Abstract. Subarctic steppe is currently restricted in interior Alaska and the Yukon Territory to steep, south-facing river bluffs. Paleoecological and biogeographic evidence suggests that some steppe taxa may have been more widespread during the Full-Glacial. We examined factors controlling the distribution of steppe taxa on an elevation gradient across a steppetundra ecotone; such analyses may help define potential Full-Glacial distributions of these taxa. Multivariate analyses suggest that species can be divided into four spatially distinct groups, but individualistic species distributions create considerable overlap among these groups. The steppe-tundra ecotone comprises a broad zone of mixing between steppe taxa and drought-tolerant alpine tundra taxa, followed by an abrupt shift to alpine shrub tundra. The transition from low steppe to tundra vegetation is primarily associated with a gradient of decreasing soil temperature. The more abrupt transition from mixed steppe-tundra to alpine shrub tundra vegetation is primarily associated with changes in soil depth and soil moisture. Variation in vegetation within steppe is associated with gradients in soil phosphorus and moisture. Greenhouse experiments on drought tolerance of two steppe and two tundra taxa suggest that the individualistic distribution of species along the ecotone is partly a function of physiological differences among species. Our analyses of vegetation-environment relationships support the hypothesis that some components of the steppe community could have been more widespread during the colder Full-Glacial.     

Vegetation of Barton Peninsula in the neighbourhood of King Sejong Station (King George Island,maritime Antarctic)

Plant communities were studied on Barton Peninsula around King Sejong Station on King George Island, maritime Antarctic. The objective of this study was to document the occurrence and distribution of plant assemblages to provide the bases for monitoring the effects of environmental changes and human impact on the vegetation of this area. Approximately 47% of the investigated area was covered by vegetation. Crustose lichens showed the highest mean cover (21%) among vegetation components. The total mean cover of the four dominant taxa, together with the other three major subdominant components, i.e., Usnea spp., Andreaea spp. and Sanionia georgico-uncinata, was 78.2% of the total cover of all the species. Lichen cover and species diversity increased with altitude and the time of exposure from snow. Lichens contributed substantially more to the increased species density and diversity than did bryophytes. Ten plant communities were recognized within the study area. All of them belong to the Antarctic cryptogam tundra formation; they were grouped into four subformations: fruticose lichen and moss cushion subformation, crustose lichen subformation, moss carpet subformation and moss hummock subformation. The moss turf subformation was not found on this region. The Antarctic herb tundra formation was also not found; however, the populations of both Antarctic vascular plants have rapidly expanded around Barton Peninsula in recent years, which may allow development of the Antarctic herb tundra formation in the future.     

Recent advance of white spruce (Picea glauca) in the coastal tundra of the eastern shore of Hudson Bay (Québec, Canada)

Aim The species‐specific response of tree‐line species to climatic forcing is a crucial topic in modelling climate‐driven ecosystem dynamics. In northern Québec, Canada, black spruce (Picea mariana) is the dominant species at the tree line, but white spruce (Picea glauca) also occurs along the maritime coast of Hudson Bay, and is expanding along the coast and on lands that have recently emerged because of isostatic uplift. Here we outline the present distribution, structure, dynamics and recent spread of white spruce from the tree line up to its northernmost position in the shrub tundra along the Hudson Bay coast. We aimed to obtain a minimum date of the arrival of the species in the area and to evaluate its dynamics relative to recent climate changes. Location White spruce populations and individuals were sampled along a latitudinal transect from the tree line to the northernmost individual in the shrub tundra along the Hudson Bay coast and in the Nastapoka archipelago in northern Québec and Nunavut, Canada (56°06′–56°32′ N). Methods White spruce populations were mapped, and the position, dimension, growth form and origin (seed or layering) of every individual recorded. Tree‐ring analyses of living and dead trees allowed an estimation of the population structure, past recruitment, growth trends and growth rate of the species. A macrofossil analysis was performed of the organic horizon of the northernmost white spruce stands and individuals. Radiocarbon dates of white spruce remains and organic matter were obtained. The rate of isostatic uplift was assessed by radiocarbon dating of drifted wood fragments. Results The first recorded establishment of white spruce was almost synchronous at all sites and occurred around ad 1660. Spruce recruitment was rather continuous at the tree line, while it showed a gap in the northern shrub tundra during the first decades of the 19th century. A vigorous, recent establishment of seedlings was observed in the shrub tundra; only wind‐exposed, low krummholz (stunted individuals) did not show any sexual regeneration. A period of suppressed growth occurred from the 1810s to the 1850s in most sites. A growth increase was evident from the second half of the 19th century and peaked in the 1880s and the 20th century. A shift from stunted to tree growth form has occurred since the mid‐19th century. No sample associated with white spruce remains gave a date older than 300 ¹⁴C years bp [calibrated age (cal.) ad 1430–1690]. Main conclusions White spruce probably arrived recently in the coastal tundra of Hudson Bay due to a delayed post‐glacial spread. The arrival of the species probably occurred during the Little Ice Age. The established individuals survived by layering during unfavourable periods, but acted as nuclei for sexual recruitment almost continuously, except in the northernmost and most exposed sites. Warmer periods were marked by strong seedling recruitment and a shift to tree growth form. Unlike white spruce, black spruce showed no evidence of an ongoing change in growth form and sexual recruitment. Ecological requirements and recent history of tree‐line species should be taken into account in order to understand the present dynamics of high‐latitude ecosystems.     

The start of the birch pollen season in Finnish Lapland: separating non‐local from local birch pollen and the implication for allergy sufferers

Determining the start of the birch pollen season requires the reliable separation of non‐local from locally produced birch pollen. The research was undertaken close to the latitudinal birch tree line at the Kevo Subarctic Research Institute (69°45′N 27°01′E) in northern Finland. By comparing phenological and aerobiological observations, the proportion of birch pollen present in the air before local anthesis commences can be delimited. We coupled this with data of pollen deposition monitored by means of a modified Tauber trap. The dominant birch species at Kevo is the mountain birch Betula pubescens ssp. czerepanovii, whereas B. pubescens ssp. pubescens is very rare, hence we consider the proportion of the southerly B. pubescens‐type pollen deposited in the pollen trap to be non‐local in origin.

We did not observe any trend towards an earlier start of the phenologically observed mountain birch anthesis at Kevo as predicted from work elsewhere. Moreover, the fixed 2.5% threshold method for determining the birch pollen season proved not to be applicable since in many years this threshold was reached before the end of continuous snow cover. The results indicate that in some years non‐local birch pollen contributes considerably to the allergen load in Lapland with up to 57% of the total birch pollen sum being recorded before the day on which local anthesis commenced, and up to 70% of the annual birch pollen deposited being of the southerly birch type.     

Beetles (Insecta,Coleoptera) in the arctic fauna: Communication 1. Faunal composition

Coleoptera, the largest insect order, appears to be subordinate to Diptera in the extent of colonization of the Arctic environment. Beetles comprise about 13% of the insect fauna of the tundra zone, yet in the high latitudes several of their families retain considerable levels of species diversity and play significant cenotic roles. The present communication reviews the circumpolar beetle fauna of the Arctic. Based on original data, literature sources and museum collections, with the use of extrapolations and analogies, the taxonomic and ecological diversity patterns of the suborders, series and families of Coleoptera are distinguished, the latitude-zonal distribution and the northernmost range limits of the species are analyzed, and their adaptations and cenotic relations are characterized.     

Seasonal patterns of life cycle of ground beetle <Emphasis Type="Italic">Carabus nitens</Emphasis> (Coleoptera,Carabidae) in southern tundra

Seasonal dynamics of activity and demographic structure of Carabus nietens in the forest-tundra and southern tundra of the Kanin Peninsula was studied. In the northernmost range, the species has a one-year life cycle with reproduction in the early season and summer larvae. C. nitens demonstrated elongated period of development from egg to imago compared to the temperate zone in the polar region. At the same time, overwintered beetles rapidly matured and the reproduction period was reduced to three decades. This synchronizes larval development and the young generation emergence. The northern populations have increased life span of imago, which leads to repeated postwinter reproduction during several seasons. As a result, the abundance of the northern C. nitens populations remains high.     

Modern pollen and stomate deposition in lake surface sediments from across the treeline on the Kola Peninsula, Russia

We sampled and analyzed surface sediments from 31 lakes along a latitudinal transect crossing the coniferous treeline on the Kola Peninsula, Russia. The major vegetation zones along the transect were tundra, birch-forest tundra, pine-forest tundra, and forest. The results indicate that the major vegetation types in our study area have distinct pollen spectra. Sum-of-squares cluster analysis and principal components analysis (PCA) groupings of pollen sites correspond to the major vegetation zones. PCA ordination of taxa indicates that the first axis separates taxa typical of the forest zone (Pinus, Picea) from taxa typical of tundra and forest-tundra zones (Polypodiaceae, Ericaceae, and Betula). The current position of the coniferous treeline, defined in our region by Pinus sylvestris, occurs roughly where Pinus pollen values reach 35% or greater. Arboreal pollen (AP)/non-arboreal pollen (NAP) ratios were calculated for each site and plotted against geographic distance along the transect. AP/NAP ratios of 7 or greater are found within pine-forest tundra and forest vegetation zones. Pinus stomates (dispersed stomatal guard cells) are absent from sites north of the coniferous treeline and all but two samples from the forested sites contain stomates. Stomate concentrations among the samples are highly variable and range from 10 to 458 per ml and positively correlate with the changing Pinus pollen values.     

Temporal and spatial distribution of microcrustacean zooplankton in relation to turbidity and other environmental factors in a large tropical lake (L. Tana, Ethiopia)

The spatial and seasonal distribution of microcrustacean zooplankton of Lake Tana (Ethiopia) was monthly studied for 2 years. Concurrently, various environmental parameters were measured and related to zooplankton distribution. Canonical Correspondence Analysis (CCA) was used to estimate the influence of abiotic factors and chlorophyll a content in structuring the zooplankton assemblage. Among the environmental factors, zooplankton abundance correlated most strongly with turbidity. Turbidity was negatively correlated with species abundance, especially for Daphnia spp. and to the least extent for Diaphanosoma spp. Analysis of variance (ANOVA) was used to determine spatial (littoral, sublittoral and pelagic zone) and temporal (four seasons) variation in zooplankton abundance. We observed significant temporal differences in zooplankton abundance, with highest densities during dry season (November–April). Only cladocerans showed significant differences in habitat use (highest densities in the sublittoral zone). %     

The effect of extreme northerly climatic conditions on the life history of the minnow, Phoxinus phoxinus (L.)

Over its exceptionally wide geographical range the minnow displays a remarkable variability in its life history. In southern England many fish mature after one year and few fish survive to their third birthday. Very few fish spawned for more than one season but a 65-mm female is estimated to spawn up to 3172 eggs over the extended spawning season. In the River Utsjoki in Finnish Lapland maturity was strongly size-dependent and delayed until the fish reached 5, 6 or even 7 years of age, with a maximum age of 13 years at a length of only 75 mm. Unlike some other small-sized species in unproductive environments, individual females continued to spawn successive clutches of eggs though over a much shorter spawning season. The maximum estimated egg production was 824 for a 65-mm fish. Temperature had a strong direct or indirect effect on growth, with significantly lower growth increments in cold summers. At a second Arctic site in eastern Lapland growth rates were higher and maturity earlier, yet clutch size was greatly reduced with a seasonal fecundity estimate of only 314. There were only small differences in size of eggs or larvae between the populations. Evidence from other species indicates that most of the observed changes over the range of the minnow are phenotypic responses to the contrasting environments. However, the different strategies displayed at the two Arctic sites could represent the results of selection for differences in pre- and post-reproductive survival.     

Distribution of water bugs (Heteroptera: Nepomorpha, Gerromorpha) over latitudinal zones in the northeast of European Russia

Data on the taxonomic composition of water bugs (Heteroptera: Nepomorpha, Gerromorpha) in the northeast of European Russia are given. A total of 34 water-bug species, which belong to 16 genera and 8 families, are recorded. The species diversity of Nepomorpha and Gerromorpha changes northwards: 33 occur in the taiga zone and only 8 species occur in the tundra zone. The bulk of the fauna consists of Eurosiberian (29%) and trans-Eurasian (26%) species; in latitude-related terms, the temperate group is most richly represented (64%). The species composition of the aquatic and semiaquatic heteropteran fauna of the region is especially similar to that of the tundra and taiga zones of Western Siberia.     

Greater shrub dominance alters breeding habitat and food resources for migratory songbirds in Alaskan arctic tundra

Climate warming is affecting the Arctic in multiple ways, including via increased dominance of deciduous shrubs. Although many studies have focused on how this vegetation shift is altering nutrient cycling and energy balance, few have explicitly considered effects on tundra fauna, such as the millions of migratory songbirds that breed in northern regions every year. To understand how increasing deciduous shrub dominance may alter breeding songbird habitat, we quantified vegetation and arthropod community characteristics in both graminoid and shrub dominated tundra. We combined measurements of preferred nest site characteristics for Lapland longspurs (Calcarius lapponicus) and Gambel's White‐crowned sparrows (Zonotrichia leucophrys gambelii) with modeled predictions for the distribution of plant community types in the Alaskan arctic foothills region for the year 2050. Lapland longspur nests were found in sedge‐dominated tussock tundra where shrub height does not exceed 20 cm, whereas White‐crowned sparrows nested only under shrubs between 20 cm and 1 m in height, with no preference for shrub species. Shrub canopies had higher canopy‐dwelling arthropod availability (i.e. small flies and spiders) but lower ground‐dwelling arthropod availability (i.e. large spiders and beetles). Since flies are the birds' preferred prey, increasing shrubs may result in a net enhancement in preferred prey availability. Acknowledging the coarse resolution of existing tundra vegetation models, we predict that by 2050 there will be a northward shift in current White‐crowned sparrow habitat range and a 20–60% increase in their preferred habitat extent, while Lapland longspur habitat extent will be equivalently reduced. Our findings can be used to make first approximations of future habitat change for species with similar nesting requirements. However, we contend that as exemplified by this study's findings, existing tundra modeling tools cannot yet simulate the fine‐scale habitat characteristics that are critical to accurately predicting future habitat extent for many wildlife species.     

Spatial structure of the soil seed bank of Poa annua L.—alien species in the Antarctica

Poa annua L. (annual bluegrass) is the only non–native flowering plant species that has successfully established a breeding population in the maritime Antarctic and has been shown to maintain a soil seed bank. The characteristic of the spatial structure of the Antarctic population of this species is the formation of distinct dense clumps—tussocks. In the temperate zone the species is only loosely tufted. We focused on the characteristics of seed deposition associated with the tussocks and some aspects of the spatial heterogeneity of the soil seed bank of P. annua in the Antarctic. We wanted to assess the microspatial structure of the soil seed bank of annual bluegrass at Arctowski Station. Therefore we compared the number of seeds deposited underneath and in the vicinity of P. annua clumps. Our results indicate that P. annua in the Antarctic maintains a soil seed bank comparable to species typical for the polar tundra. The microspatial structure of P. annua soil seed bank in the Antarctic is highly associated with the presence of tussocks. Seeds are deposited underneath the tussock rather than in the vicinity of the clump. Our results also indicate that seeds are able to survive the Antarctic winter and readily germinate under optimal conditions.     

Decomposition of mountain birch leaf litter at the forest-tundra ecotone in the Fennoscandian mountains in relation to climate and soil conditions

Litter decomposition is a key process in terrestrial ecosystems, releasing nutrients, returning CO₂ to the atmosphere, and contributing to the formation of humus. Litter decomposition is strongly controlled both by climate and by litter quality: global warming scenarios involving shifts in vegetation communities are therefore of particular interest in this context. The objective of the present study was to quantify the role of climatic environment and underlying substrate chemistry for the decomposition of standard mountain birch (Betula pubescens Ehrh. spp. czerepanovii) leaf litter at four sites, spanning the forest-tundra ecotone, in the Fennoscandian mountain range. Litter quality effects were thus held constant, but the study incorporated systematic changes in (i) latitude/altitude, (ii) `continentality', and (iii) vegetation community at each site, together with (iv) experimental manipulation of temperature using passive warming systems. The study was undertaken during a 3 year period, and forms part of a broader investigation of forest-tundra ecotone dynamics in the Fennoscandian mountains. Our results showed (1) higher decomposition rates in forest sites compared to tundra, (2) that the difference between the two vegetation communities was most pronounced at the more maritime sites, and (3) that chemistry of litter remaining after the three years experiment varied according to site and vegetation community (e.g. at the most southerly site, more lignin had decomposed at tundra communities compared with the forest). (4) Surface temperature explained 58% of the variation in mass loss at forest sites; at tundra sites, however, we hypothesise that litter moisture content was the more important factor. (5) Experimental warming lent weight to this hypothesis by reducing rates of mass loss: this reduction was likely the result of surface soil drying, an artefact of the warming treatment. We conclude that a replacement of tundra by forest would likely accelerate litter decomposition both via changes in surface and near-surface temperature and moisture regimes, although the strength of this response will vary between maritime and continental parts of the mountain range.     

Population structure, recruitment, and succession of the brown alga, Padina boryana Thivy (Dictyotales, Heterokontophyta), at an exposed shore of Sirinart National Park and a sheltered area of Tang Khen Bay, Phuket Province, Thailand

The brown algal genus, Padina, has a worldwide distribution in tropical and subtropical climate zones. Padina individuals are common and sometimes dominant in both the intertidal and shallow subtidal regions associated with coral reefs. We investigated the population structure and recruitment of two populations of Padina boryana at an exposed shore in Sirinart National Park (SNP) and a sheltered shore of Tang Khen Bay (TKB), Phuket Province, Thailand. From September 2005 to August 2006, the number of released spores and the height and radius of fronds were measured monthly. New recruits were recorded and monitored on manipulated permanent plots 0.25 m². The experiment was carried out monthly over a year. We found Padina recruits 1 month after the plots were cleared at both sites. There was a significant difference in percentage cover by new individuals between the two locations (P < 0.05) and also at the shore levels within the sites. At SNP, the highest recruitment was found in the uppermost zone, while at TKB high recruitment occurred at all shore levels except at 80-100 m from the shore. The factors influencing recruitment of P. boryana include wave motion, competition with other earlier successional species and sediment. The high availability of reproductive spores throughout the year, the successful recruitment and the Dictyerpa stage promote the successful establishment of Padina populations.     

Butterflies (Lepidoptera, Rhopalocera) in the Arctic fauna

The Arctic fauna includes 106 species of diurnal butterflies: Papilionidae (6 species), Pieridae (20), Lycaenidae (18), Nymphalidae (30), Satyridae (27), and Hesperiidae (5). Among them, representatives of the family Nymphalidae predominate as to the features characterizing the biological progress in the Arctic, as well as to the number of the most strongly pronounced arctic forms. The family Satyridae shares the first place with Nymphalidae by the number of species, but differs from the latter in the uneven or local distribution. The family Pieridae demonstrates a wide distribution of polyzonal and boreal species in the tundra zone. The distribution patterns of Lycaenidae are different in the Eurasian and Beringian-American sectors. Species of Papilionidae and Hesperiidae occur only in the southern part of the tundra zone. Each family is characterized by specific distribution in the Arctic subzones and landscapes and by latitudinal trends in its specific ratio in the faunas. There are 30 to 40 arctic species, including arctic proper (euarctic and hemiarctic) and hypoarctic, arctoalpine, arctomontane, and arctoboreal species. The species developing successfully under high-latitude conditions are Boloria chariclea, B. polaris, B. improba, Colias nastes, C. hecla, and Erebia fasciata; the first two species can be considered true euarctic forms. Specific features of the latitudinal and longitudinal distribution of the butterfly species in different parts of the Arctic are discussed.     

Where do the treeless tundra areas of northern highlands fit in the global biome system: toward an ecologically natural subdivision of the tundra biome

According to some treatises, arctic and alpine sub‐biomes are ecologically similar, whereas others find them highly dissimilar. Most peculiarly, large areas of northern tundra highlands fall outside of the two recent subdivisions of the tundra biome. We seek an ecologically natural resolution to this long‐standing and far‐reaching problem. We studied broad‐scale patterns in climate and vegetation along the gradient from Siberian tundra via northernmost Fennoscandia to the alpine habitats of European middle‐latitude mountains, as well as explored those patterns within Fennoscandian tundra based on climate–vegetation patterns obtained from a fine‐scale vegetation map. Our analyses reveal that ecologically meaningful January–February snow and thermal conditions differ between different types of tundra. High precipitation and mild winter temperatures prevail on middle‐latitude mountains, low precipitation and usually cold winters prevail on high‐latitude tundra, and Scandinavian mountains show intermediate conditions. Similarly, heath‐like plant communities differ clearly between middle latitude mountains (alpine) and high‐latitude tundra vegetation, including its altitudinal extension on Scandinavian mountains. Conversely, high abundance of snowbeds and large differences in the composition of dwarf shrub heaths distinguish the Scandinavian mountain tundra from its counterparts in Russia and the north Fennoscandian inland. The European tundra areas fall into three ecologically rather homogeneous categories: the arctic tundra, the oroarctic tundra of northern heights and mountains, and the genuinely alpine tundra of middle‐latitude mountains. Attempts to divide the tundra into two sub‐biomes have resulted in major discrepancies and confusions, as the oroarctic areas are included in the arctic tundra in some biogeographic maps and in the alpine tundra in others. Our analyses based on climate and vegetation criteria thus seem to resolve the long‐standing biome delimitation problem, help in consistent characterization of research sites, and create a basis for further biogeographic and ecological research in global tundra environments.     

Fast assimilate turnover revealed by in situ 13CO2 pulse-labelling in Subarctic tundra

Climatic changes in Arctic regions are likely to have significant impacts on vegetation composition and physiological responses of different plant types, with implications for the regional carbon (C) cycle. Here, we explore differences in allocation and turnover of assimilated C in two Subarctic tundra communities. We used an in situ ¹³C pulse at mid-summer in Swedish Lapland to investigate C allocation and turnover in four contrasting tundra plant communities. We found a high rate of turnover of assimilated C in leaf tissues of Betula nana and graminoid vegetation at the height of the growing season, with a mean residence time of pulse-derived ¹³C of 1.1 and 0.7?days, respectively. One week after the pulse, c. 20 and 15%, respectively, of assimilated label-C remained in leaf biomass, representing most likely allocation to structural biomass. For the perennial leaf tissue of the graminoid communities, a remainder of approximately 5% of the pulse-derived C was still traceable after 1?year, whereas none was detectable in Betula foliage. The results indicate a relatively fast C turnover and small belowground allocation during the active growing season of recent assimilates in graminoid communities, with comparatively slower turnover and greater investment in belowground allocation by B. nana vegetation.